On the Origin of Species by Means of Natural Selection : Chapter III-A

Bears on natural selection -- The term used in a wide sense -- Geometrical
powers of increase -- Rapid increase of naturalised animals and plants --
Nature of the checks to increase -- Competition universal -- Effects of
climate -- Protection from the number of individuals -- Complex relations
of all animals and plants throughout nature -- Struggle for life most
severe between individuals and varieties of the same species; often severe
between species of the same genus -- The relation of organism to organism
the most important of all relations.

Before entering on the subject of this chapter, I must make a few
preliminary remarks, to show how the struggle for existence bears on
Natural Selection. It has been seen in the last chapter that amongst
organic beings in a state of nature there is some individual variability;
indeed I am not aware that this has ever been disputed. It is immaterial
for us whether a multitude of doubtful forms be called species or
sub-species or varieties; what rank, for instance, the two or three hundred
doubtful forms of British plants are entitled to hold, if the existence of
any well-marked varieties be admitted. But the mere existence of
individual variability and of some few well-marked varieties, though
necessary as the foundation for the work, helps us but little in
understanding how species arise in nature. How have all those exquisite
adaptations of one part of the organisation to another part, and to the
conditions of life, and of one distinct organic being to another being,
been perfected? We see these beautiful co-adaptations most plainly in the
woodpecker and missletoe; and only a little less plainly in the humblest
parasite which clings to the hairs of a quadruped or feathers of a bird; in
the structure of the beetle which dives through the water; in the plumed
seed which is wafted by the gentlest breeze; in short, we see beautiful
adaptations everywhere and in every part of the organic world.

Again, it may be asked, how is it that varieties, which I have called
incipient species, become ultimately converted into good and distinct
species, which in most cases obviously differ from each other far more than
do the varieties of the same species? How do those groups of species,
which constitute what are called distinct genera, and which differ from
each other more than do the species of the same genus, arise? All these
results, as we shall more fully see in the next chapter, follow inevitably
from the struggle for life. Owing to this struggle for life, any
variation, however slight and from whatever cause proceeding, if it be in
any degree profitable to an individual of any species, in its infinitely
complex relations to other organic beings and to external nature, will tend
to the preservation of that individual, and will generally be inherited by
its offspring. The offspring, also, will thus have a better chance of
surviving, for, of the many individuals of any species which are
periodically born, but a small number can survive. I have called this
principle, by which each slight variation, if useful, is preserved, by the
term of Natural Selection, in order to mark its relation to man's power of
selection. We have seen that man by selection can certainly produce great
results, and can adapt organic beings to his own uses, through the
accumulation of slight but useful variations, given to him by the hand of
Nature. But Natural Selection, as we shall hereafter see, is a power
incessantly ready for action, and is as immeasurably superior to man's
feeble efforts, as the works of Nature are to those of Art.

We will now discuss in a little more detail the struggle for existence. In
my future work this subject shall be treated, as it well deserves, at much
greater length. The elder De Candolle and Lyell have largely and
philosophically shown that all organic beings are exposed to severe
competition. In regard to plants, no one has treated this subject with
more spirit and ability than W. Herbert, Dean of Manchester, evidently the
result of his great horticultural knowledge. Nothing is easier than to
admit in words the truth of the universal struggle for life, or more
difficult--at least I have found it so--than constantly to bear this
conclusion in mind. Yet unless it be thoroughly engrained in the mind, I
am convinced that the whole economy of nature, with every fact on
distribution, rarity, abundance, extinction, and variation, will be dimly
seen or quite misunderstood. We behold the face of nature bright with
gladness, we often see superabundance of food; we do not see, or we forget,
that the birds which are idly singing round us mostly live on insects or
seeds, and are thus constantly destroying life; or we forget how largely
these songsters, or their eggs, or their nestlings, are destroyed by birds
and beasts of prey; we do not always bear in mind, that though food may be
now superabundant, it is not so at all seasons of each recurring year.

I should premise that I use the term Struggle for Existence in a large and
metaphorical sense, including dependence of one being on another, and
including (which is more important) not only the life of the individual,
but success in leaving progeny. Two canine animals in a time of dearth,
may be truly said to struggle with each other which shall get food and
live. But a plant on the edge of a desert is said to struggle for life
against the drought, though more properly it should be said to be dependent
on the moisture. A plant which annually produces a thousand seeds, of
which on an average only one comes to maturity, may be more truly said to
struggle with the plants of the same and other kinds which already clothe
the ground. The missletoe is dependent on the apple and a few other trees,
but can only in a far-fetched sense be said to struggle with these trees,
for if too many of these parasites grow on the same tree, it will languish
and die. But several seedling missletoes, growing close together on the
same branch, may more truly be said to struggle with each other. As the
missletoe is disseminated by birds, its existence depends on birds; and it
may metaphorically be said to struggle with other fruit-bearing plants, in
order to tempt birds to devour and thus disseminate its seeds rather than
those of other plants. In these several senses, which pass into each
other, I use for convenience sake the general term of struggle for
existence.

A struggle for existence inevitably follows from the high rate at which all
organic beings tend to increase. Every being, which during its natural
lifetime produces several eggs or seeds, must suffer destruction during
some period of its life, and during some season or occasional year,
otherwise, on the principle of geometrical increase, its numbers would
quickly become so inordinately great that no country could support the
product. Hence, as more individuals are produced than can possibly
survive, there must in every case be a struggle for existence, either one
individual with another of the same species, or with the individuals of
distinct species, or with the physical conditions of life. It is the
doctrine of Malthus applied with manifold force to the whole animal and
vegetable kingdoms; for in this case there can be no artificial increase of
food, and no prudential restraint from marriage. Although some species may
be now increasing, more or less rapidly, in numbers, all cannot do so, for
the world would not hold them.

There is no exception to the rule that every organic being naturally
increases at so high a rate, that if not destroyed, the earth would soon be
covered by the progeny of a single pair. Even slow-breeding man has
doubled in twenty-five years, and at this rate, in a few thousand years,
there would literally not be standing room for his progeny. Linnaeus has
calculated that if an annual plant produced only two seeds--and there is no
plant so unproductive as this--and their seedlings next year produced two,
and so on, then in twenty years there would be a million plants. The
elephant is reckoned to be the slowest breeder of all known animals, and I
have taken some pains to estimate its probable minimum rate of natural
increase: it will be under the mark to assume that it breeds when thirty
years old, and goes on breeding till ninety years old, bringing forth three
pair of young in this interval; if this be so, at the end of the fifth
century there would be alive fifteen million elephants, descended from the
first pair.

But we have better evidence on this subject than mere theoretical
calculations, namely, the numerous recorded cases of the astonishingly
rapid increase of various animals in a state of nature, when circumstances
have been favourable to them during two or three following seasons. Still
more striking is the evidence from our domestic animals of many kinds which
have run wild in several parts of the world: if the statements of the rate
of increase of slow-breeding cattle and horses in South America, and
latterly in Australia, had not been well authenticated, they would have
been quite incredible. So it is with plants: cases could be given of
introduced plants which have become common throughout whole islands in a
period of less than ten years. Several of the plants now most numerous
over the wide plains of La Plata, clothing square leagues of surface almost
to the exclusion of all other plants, have been introduced from Europe; and
there are plants which now range in India, as I hear from Dr. Falconer,
from Cape Comorin to the Himalaya, which have been imported from America
since its discovery. In such cases, and endless instances could be given,
no one supposes that the fertility of these animals or plants has been
suddenly and temporarily increased in any sensible degree. The obvious
explanation is that the conditions of life have been very favourable, and
that there has consequently been less destruction of the old and young, and
that nearly all the young have been enabled to breed. In such cases the
geometrical ratio of increase, the result of which never fails to be
surprising, simply explains the extraordinarily rapid increase and wide
diffusion of naturalised productions in their new homes.

In a state of nature almost every plant produces seed, and amongst animals
there are very few which do not annually pair. Hence we may confidently
assert, that all plants and animals are tending to increase at a
geometrical ratio, that all would most rapidly stock every station in which
they could any how exist, and that the geometrical tendency to increase
must be checked by destruction at some period of life. Our familiarity
with the larger domestic animals tends, I think, to mislead us: we see no
great destruction falling on them, and we forget that thousands are
annually slaughtered for food, and that in a state of nature an equal
number would have somehow to be disposed of.

The only difference between organisms which annually produce eggs or seeds
by the thousand, and those which produce extremely few, is, that the
slow-breeders would require a few more years to people, under favourable
conditions, a whole district, let it be ever so large. The condor lays a
couple of eggs and the ostrich a score, and yet in the same country the
condor may be the more numerous of the two: the Fulmar petrel lays but one
egg, yet it is believed to be the most numerous bird in the world. One fly
deposits hundreds of eggs, and another, like the hippobosca, a single one;
but this difference does not determine how many individuals of the two
species can be supported in a district. A large number of eggs is of some
importance to those species, which depend on a rapidly fluctuating amount
of food, for it allows them rapidly to increase in number. But the real
importance of a large number of eggs or seeds is to make up for much
destruction at some period of life; and this period in the great majority
of cases is an early one. If an animal can in any way protect its own eggs
or young, a small number may be produced, and yet the average stock be
fully kept up; but if many eggs or young are destroyed, many must be
produced, or the species will become extinct. It would suffice to keep up
the full number of a tree, which lived on an average for a thousand years,
if a single seed were produced once in a thousand years, supposing that
this seed were never destroyed, and could be ensured to germinate in a
fitting place. So that in all cases, the average number of any animal or
plant depends only indirectly on the number of its eggs or seeds.

In looking at Nature, it is most necessary to keep the foregoing
considerations always in mind--never to forget that every single organic
being around us may be said to be striving to the utmost to increase in
numbers; that each lives by a struggle at some period of its life; that
heavy destruction inevitably falls either on the young or old, during each
generation or at recurrent intervals. Lighten any check, mitigate the
destruction ever so little, and the number of the species will almost
instantaneously increase to any amount. The face of Nature may be compared
to a yielding surface, with ten thousand sharp wedges packed close together
and driven inwards by incessant blows, sometimes one wedge being struck,
and then another with greater force.

What checks the natural tendency of each species to increase in number is
most obscure. Look at the most vigorous species; by as much as it swarms
in numbers, by so much will its tendency to increase be still further
increased. We know not exactly what the checks are in even one single
instance. Nor will this surprise any one who reflects how ignorant we are
on this head, even in regard to mankind, so incomparably better known than
any other animal. This subject has been ably treated by several authors,
and I shall, in my future work, discuss some of the checks at considerable
length, more especially in regard to the feral animals of South America.
Here I will make only a few remarks, just to recall to the reader's mind
some of the chief points. Eggs or very young animals seem generally to
suffer most, but this is not invariably the case. With plants there is a
vast destruction of seeds, but, from some observations which I have made, I
believe that it is the seedlings which suffer most from germinating in
ground already thickly stocked with other plants. Seedlings, also, are
destroyed in vast numbers by various enemies; for instance, on a piece of
ground three feet long and two wide, dug and cleared, and where there could
be no choking from other plants, I marked all the seedlings of our native
weeds as they came up, and out of the 357 no less than 295 were destroyed,
chiefly by slugs and insects. If turf which has long been mown, and the
case would be the same with turf closely browsed by quadrupeds, be let to
grow, the more vigorous plants gradually kill the less vigorous, though
fully grown, plants: thus out of twenty species growing on a little plot
of turf (three feet by four) nine species perished from the other species
being allowed to grow up freely.

The amount of food for each species of course gives the extreme limit to
which each can increase; but very frequently it is not the obtaining food,
but the serving as prey to other animals, which determines the average
numbers of a species. Thus, there seems to be little doubt that the stock
of partridges, grouse, and hares on any large estate depends chiefly on the
destruction of vermin. If not one head of game were shot during the next
twenty years in England, and, at the same time, if no vermin were
destroyed, there would, in all probability, be less game than at present,
although hundreds of thousands of game animals are now annually killed. On
the other hand, in some cases, as with the elephant and rhinoceros, none
are destroyed by beasts of prey: even the tiger in India most rarely dares
to attack a young elephant protected by its dam.

Climate plays an important part in determining the average numbers of a
species, and periodical seasons of extreme cold or drought, I believe to be
the most effective of all checks. I estimated that the winter of 1854-55
destroyed four-fifths of the birds in my own grounds; and this is a
tremendous destruction, when we remember that ten per cent. is an
extraordinarily severe mortality from epidemics with man. The action of
climate seems at first sight to be quite independent of the struggle for
existence; but in so far as climate chiefly acts in reducing food, it
brings on the most severe struggle between the individuals, whether of the
same or of distinct species, which subsist on the same kind of food. Even
when climate, for instance extreme cold, acts directly, it will be the
least vigorous, or those which have got least food through the advancing
winter, which will suffer most. When we travel from south to north, or
from a damp region to a dry, we invariably see some species gradually
getting rarer and rarer, and finally disappearing; and the change of
climate being conspicuous, we are tempted to attribute the whole effect to
its direct action. But this is a very false view: we forget that each
species, even where it most abounds, is constantly suffering enormous
destruction at some period of its life, from enemies or from competitors
for the same place and food; and if these enemies or competitors be in the
least degree favoured by any slight change of climate, they will increase
in numbers, and, as each area is already fully stocked with inhabitants,
the other species will decrease. When we travel southward and see a
species decreasing in numbers, we may feel sure that the cause lies quite
as much in other species being favoured, as in this one being hurt. So it
is when we travel northward, but in a somewhat lesser degree, for the
number of species of all kinds, and therefore of competitors, decreases
northwards; hence in going northward, or in ascending a mountain, we far
oftener meet with stunted forms, due to the directly injurious action of
climate, than we do in proceeding southwards or in descending a mountain.
When we reach the Arctic regions, or snow-capped summits, or absolute
deserts, the struggle for life is almost exclusively with the elements.